Ameiva Auberi Ustulata, Squamata

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WWW.IRCF.ORG/REPTILESANDAMPHIBIANSJOURNALTABLE OF CONTENTS IRCF REPTILES & AMPHIBIANS IRCF REPTILES • VOL15, & N AMPHIBIANSO 4 • DEC 2008 189• 19(2):85–89 • JUNE 2012

IRCF REPTILES & AMPHIBIANS CONSERVATION AND NATURAL HISTORY

TABLE OF CONTENTS

FEATURE ARTICLES Courtship. Chasing Bullsnakes (Pituophis Behavior catenifer sayi) in Wisconsin: in the Cuban Ameiva On the Road to Understanding the Ecology and Conservation of the Midwest’s Giant Serpent ...... Joshua M. Kapfer 190 . The Shared History of Treeboas (Corallus grenadensis) and Humans on Grenada: (AmeivaA Hypothetical auberi Excursion ...... ustulata, Squamata:Robert W. Henderson Teiidae) 198 fromRESEARCH the ARTICLES Siboney-Juticí Ecological Reserve . The Texas Horned Lizard in Central and Western Texas ...... Emily Henry, Jason Brewer, Krista Mougey, and Gad Perry 204 . The Knight Anole (Anolis equestris) in Florida ...... Brianin J. Camposano, Eastern Kenneth L. Krysko, Kevin M.Cuba Enge, Ellen M. Donlan, and Michael Granatosky 212 CONSERVATION ALERT Yasel U. Alfonso1 and Javier Torres2 . World’s Mammals in Crisis ...... 220 1 . More Than MammalsCentro de ...... Aplicaciones Tecnológicas para el Desarrollo Sostenible (CATEDES/CITMA) I, 223 . The “DowAgramonte Jones Index” No. 818of Biodiversity e/ Aguilera ...... y Prado, Guantánamo, CP 95100, Cuba ([email protected]) 225 2Departamento de Biología Animal y Humana, Facultad de Biología, Universidad de la Habana, HUSBANDRYCalle 25 # 455 e/ J e I, Vedado, Plaza de la Revolución, La Habana, CP 10400, Cuba ([email protected]) . Captive Care of the Central Netted Dragon ...... Shannon Plummer 226

PROFILE . Kraig Adler: A Lifetime Promoting Herpetology ...... Michael L. Treglia 234

Abstract.—COMMENTARYThe genus Ameiva Meyer 1795 comprises 23 species of which 14 are found in Central and South America and. 18The Turtlesoccur Have in Been the Watching West MeIndies...... These lizards occupy diverse habitats (e.g., Eric grasslands, Gangloff 238 tropical forests, sandy beaches) but most species appear to prefer open areas. They are ground-dwellers and active diurnal BOOK REVIEW foragers. Cuba harbors only one species, . However, 40 subspecies are distributed widely across . Threatened Amphibians of the World editedAmeiva by S.N. Stuart,auberi M. Hoffmann, J.S. Chanson, N.A. Cox, the main island,R. Berridge,adjacent P. Ramani, cays, and and B.E. Youngon into ...... the Bahamas. Reproductive behavior hasRobert beenPowell documented243 in some species in the family Teiidae, but not for A. auberi, about which only one published account reported the underground CONSERVATION deposition RESEARCH of a single REPORTS: egg. Herein, Summaries of we Published describe Conservation for Researchthe first Reports time ...... the reproductive 245 behavior  NATURAL HISTORY RESEARCH REPORTS: Summaries of Published Reports on Natural History ...... 247 (except ovipositioning) NEWBRIEFS ...... of A. auberi ustulata in natural habitat in the Ecological Reserve Siboney-Juticí 248 (eastern Cuba) on 22–25EDITORIAL July INFORMATION2009. Between ...... 1200 and 1600 h, we observed six mating pairs directly 251 or while using  FOCUS ON CONSERVATION: A Project You Can Support ...... 252 monocular Nikon Fieldscopes. We describe three sequential stages of reproductive behavior: (1) Pursuit, (2) premating and mating, and (3) excavation, with a total duration of 64.8 ± 17.6 min (46–92 min).

he family Teiidae Gray 1827,Front with Cover. a distribution Shannon Plummer. lim- SmithBack 1968), Cover. Michael A. festiva Kern (Smith 1968), A. fuscata (Somma and Tited to the Western Hemisphere,Totat etcontains velleseque audantnine mo genera BrooksTotat et 1976),velleseque audantA. pleii mo (Censky 1995a, 1995b), and A. exsul estibus inveliquo velique rerchil estibus inveliquo velique rerchil (Hower and Hedges 2003). One oferspienimus, these, the quos genus accullabo. Ameiva Ilibus erspienimus,(Lewis 1986, quos accullabo. Rodríguez-Ramirez Ilibus and Lewis 1991, Lewis et al. Meyer 1795, includes 32 species, of whichaut dolor 14apicto occur invere inpe dolumCentral 2000).aut dolor apicto invere pe dolum fugiatis maionsequat eumque fugiatis maionsequat eumque and South America and the remainingmoditia 18 erere in thenonsedis West ma sectiatur Indies. moditia The erere Cuban nonsedis ma Ameiva sectia- (Ameiva auberi Cocteau 1838) con- These species occupy diverse habitats,ma derrovitae including voluptam, grasslands, as quos tainstur ma 40 derrovitae currently voluptam, recognized as subspecies. Twenty-eight of tropical forests, and sandy beaches (Schwartzaccullabo. and Henderson these are widely distributed across the Cuban archipelago, 1991, Hower and Hedges 2003, Henderson and Powell including a number of tiny cays (Gali and Garrido 1986, 2009), where most West Indian species appear to prefer open Rodríguez 2000). The only aspect of the species’ reproductive areas. These lizards are generally characterized as terrestrial biology to have been described is a single instance of ovide- diurnally active foragers (e.g., Schwartz and Henderson 1991, position (Estrada et al. 1987). Herein we describe for the first Sproston et al. 1999, Henderson and Powell 2009), although time courtship behavior in six mating pairs of Ameiva auberi a few individuals of some species have been observed ascend- ustulata Schwartz 1970 in their natural habitat. ing tree trunks in search of prey (Powell and Censky 2002). Despite the diversity and abundance of these lizards little Materials and Methods is known about their natural history, particularly reproduc- We conducted fieldwork on 22–25 July 2009 in the tion. Reproductive behavior has been described in other Ecological Reserve Siboney-Juticí, Santiago de Cuba (Fig. 1), species in the genus Ameiva, including A. ameiva (Simmons where Ameiva auberi ustulata is abundant (Fong et al. 2005). 1975, Quesnel 1978, Costa et al. 2010), A. chrysolaema Pairs were observed directly or using a monocular Nikon (Noble and Teale 1930), A. quadrilineata (Hirth 1963, Fieldscope. Observations occurred between 1200 and 1600 h

Figure 1. Locations of observations of mating Ameiva auberi ustulata in the Ecological Reserve Siboney-Juticí, Santiago de Cuba, Cuba. and were made from distances of 5.7–21.4 m (15.8 ± 5.8 m). side opposite the bitten limb, in effect forming a horseshoe, We measured the duration of each stage with a digital chro- the ends of which are the head and vent of the male (Fig. 2B). nometer (rounded to the nearest 0.1 s). Means are presented Subsequent separation (Fig. 2C) is quick and abrupt, with ± one standard deviation. individuals often separating by >50 cm. Excavation lasted 26.5 ± 17.4 min (12.7–53.4 min) and Results entailed both individuals digging a hole with a diameter of We observed six copulating pairs, for which we described ~2 cm and a depth to as much as 28 cm (Figs. 2D–E). Both three behavioral stages: (1) pursuit, (2) premating and mat- lizards repeatedly entered and left the hole until excavation ing, and (3) excavation. Pursuit lasted 32.0 ± 4.0 min (27.6– was concluded (Fig. 2F). Entire episodes lasted 64.8 ± 17.6 37.4 min). Males actively pursue females and might engage min (46–92 min). with other males competing for the same female. Some male aggression might be directed at the potential mate. During Discussion these engagements, a large and apparently dominant male All of our observations of mating occurred from 1200–1600 bites or attacks other nearby males, often chasing them con- h. Sampedro et al. (1982) indicated that the activity period siderable distances before returning to the female. Once for this species is 900–1600 h, with a peak at 1100–1200 h. other males are dispersed, the winner initiates actual court- Activity periods for other West Indian species of Ameiva are ship by closely following the female along a serpentine trajec- unimodal or bimodal, usually with peaks in both instances at tory (Figs. 2A & 3C), often panting (more frequent ventila- mid- to late morning (Henderson and Powell 2009). tion than usual with evident expansion of the gular region). These observations of mating behavior in A. auberi ustu- Tongue-flicking rates increase in both individuals, with the lata in late July might be timed so hatchlings emerge during male frequently licking the back and sides of the female. the late-summer/fall rainy season, when resources presum- The second stage (premating and mating) lasted 3.5 ± ably are most abundant. Although not known for Cuban 1.1 min (2.3–5.5 min). Males immobilize a female by biting A. auberi, Smith’s (1968) study of reproductive activity in a hindlimb and twisting until achieving penetration from the Costa Rican A. festiva and A. quadrilineata indicated that ovi-

86 ALFONSO AND TORRES IRCF REPTILES & AMPHIBIANS • 19(2):85–89 • JUNE 2012

B C

D E

Figure 2. Courtship behavior in Ameiva auberi ustulata: A. premating, the male is in a superior position; B. copulation; C. separation; D–E. excavation; F. the female is in the hole the male is entering. 87 ALFONSO AND TORRES IRCF REPTILES & AMPHIBIANS • 19(2):85–89 • JUNE 2012 positioning occurs 17–21 days after ovulation. We did not observe hatchlings nor see any signs of actual ovipositioning during the sampling period. Copulating A. ameiva tobagana on Grenada (Simmons et al. 2005) and A. erythrocephala on St. Eustatius (Kerr et al. 2005) were observed in June. Meier et al. (1993) observed A. polops on Green Cay (St. Croix) mating in August. Quesnel (1978) described a four-stage courtship based on observation of 19 pairs of A. ameiva tobagana in their natural habitat. The first three stages (summarized in Fig. 3A) involve male behavior prior to mating. During this time, a male cir- cles a female that occasionally moves more slowly to allow the male to approach. These circular movements are repeated with increasingly smaller diameters until mating occurs. For A. auberi ustulata, we combine all pre-mating behaviors into a single category (pursuit). Pairs do not engage in the circular movements seen in A. ameiva tobagana. Instead, male movements parallel those of the female and tend to follow a serpentine path (Fig. 3C). The fourth stage described by Quesnel (1978) corre- sponds to the second stage we describe in the courtship of A. Figure 4. Macroscopic aspect of the reproductive systems in (A) a mature auberi ustulata. However, relative position and posture vary. male (SVL = 62.5 mm) and (B) a mature female (SVL = 60.8 mm) Ameiva auberi hardyi in the Parque Nacional Desembarco del Granma, Granma, In both instances, males are on top of the females, but male A. Cuba. White bar = 5 mm. ameiva tobagana are diagonal or almost perpendicular relative to the female (Fig. 3B), whereas male A. auberi ustulata curl around the female (Fig. 3D), similar to what Noble and Teale ductive success in many instances offset the costs of increased (1930) described for A. chrysolaema. Copulatory duration of exposure to predators. the two species was almost identical (2.01 ± 0.3 min in A. The third stage (excavation) has not been reported for ameiva tobagana and 2.03 ± 0.25 min in A. auberi ustulata). other species of Ameiva or for other subspecies of A. auberi. This relatively short time might serve to minimize exposure At this time, females are not gravid or ready to lay eggs, but to predators at a time when lizards are particularly vulnerable. the hole might be used for ovipositioning at a later date if ovi- Lima and Dill (1990) noted that reproductive activities can positioning behavior of A. auberi ustulata is similar to that of expose both sexes of certain taxa to increased risk of preda- Saw-scaled Curlytails (Leiocephalus carinatus: Leiocephalidae), tion, but Cooper (1999) indicated that the benefits of repro- another diurnally active terrestrial lizard known to occur in the Ecological Reserve Siboney-Juticí. Eggs of the latter species are deposited in holes dug previously and carefully covered and camouflaged with dirt and surface debris (unpubl. data). The only data on eggs of A. auberi (Estrada et al. 1987) are from a single white, ovoid egg buried in a semideciduous forest over karst on the Peninsula of Guanacahabibes (western Cuba). That egg measured 11.8 x 15.8 mm, but recent (August 2011) observations in another subspecies from southeastern Cuba (A. auberi hardyi) revealed two oviductal eggs that averaged 24.2 mm in length (Fig. 4). Additional oviductal eggs in earlier stages of development were also evident (unpubl. data). Even if clutch size is only one or two, females might lay multiple clutches during a reproductive season, as in A. exsul (Rodríguez-Ramirez and Lewis 1991) and A. pleii (Censky 1995b). Clutch size in A. fuscata is 5 (N = 1; A. James in Henderson and Powell 2009), whereas clutch size is 1–7 in Anguillian A. pleii (Censky 1995a) and in Puerto Rican Figure 3. Courtship (A) and mating (B) in Ameiva ameiva tobagana (from Quesnel 1978); pursuit (C) and copulation (D) in A. auberi ustulata (this A. exsul (Wolcott 1923, Lewis 1986, Rodríguez-Ramirez and study). Lewis 1991) and A. wetmorei (Rodríguez-Ramirez and Lewis 88 ALFONSO AND TORRES IRCF REPTILES & AMPHIBIANS • 19(2):85–89 • JUNE 2012

1991), with eggs measuring 20–22 x 13–15.5 mm in the lat- a severely altered habitat. Caribbean Journal of Science 41:162–169. ter species (Wolcott 1923). Lewis, A.R. 1986. Body size and growth in two populations of the Puerto Rican Ground Lizard (Teiidae). Journal of Herpetology 20:190–195. Lewis, A.R., G. Tirano, and J. Sepulveda. 2000. Body size and paternity in a teiid Acknowledgments lizard (Ameiva exsul). Journal of Herpetology 34:110–120. We thank the personnel of the biological station at the Lima, S.L. and L.M. Dill. 1990. Behavioral decisions made under the risk of pre- Siboney-Juticí Ecological Reserve for their hospitality and dation: A review and prospectus. Canadian Journal of Zoology 68:619–640. assistance. Dr. O.H. Garrido provided helpful comments on Meier, A.J., R.E. Noble, and S.L. Rathbun. 1993. Population status and notes on the biology of the St. Croix Ground Lizard on Green Cay (St. Croix, U.S. an earlier draft of this manuscript. Robert Powell provided Virgin Islands). Caribbean Journal of Science 29:147–152. helpful literature, comments, and suggestions on an earlier Noble, G.K. and H.K. Teale. 1930. The courtship of some iguanid and teiid liz- draft of this manuscript. ards. Copeia 1930:54–56. Powell, R and E.J. Censky. 2002. Ameiva alboguttata. Arboreal activity. Herpetological Review 33:50. Literature Cited Quesnel, V.C. 1978. The reproductive behaviour of the lizard, Ameiva ameiva Censky, E.J. 1995a. Reproduction in two Lesser Antillean populations of Ameiva tobagana. Living World 1978–1979:16–18. plei (Teiidae). Journal of Herpetology 29:553–560. Rodríguez-Ramirez, J. and A.R. Lewis. 1991. Reproduction in the Puerto Rican Censky, E.J. 1995b. Mating strategy and reproductive success in the teiid lizard, teiids Ameiva exsul and A. wetmorei. Herpetologica 47:395–403. Ameiva plei. Behaviour 132:529–557. Rodríguez-Schettino, L. 2000. Cuban reptiles: Original citations, holotypes and Cooper, W.E., Jr. 1999. Tradeoffs between courtship, fighting, and antipreda- geographic range. Smithsonian Herpetological Information. Service No. 125. tory behavior by a lizard, . Eumeces laticeps Behavioral Ecology and Sociobiology Smithsonian Institution, Washington, D.C. 47:54–59. Sampedro, A., V. Berovides, and O. Torres. 1982. Morfometría, alimentación, y Costa, H.C., E.T. da Silva, P.S. Campos, M.P. da Cunha Oliveira, A.V. Nuses, actividad de Ameiva auberi (Reptilia: Teiidae) en el sur de la región Oriental and P. da Silva Santos. 2010. The corpse bride: A case of Davian behaviour in de Cuba. Ciencias Biológicas 7:105–112. the Green Ameiva (Ameiva ameiva) in southeastern Brazil. Herpetology Notes 3:79–83. Schwartz, A. and R.W. Henderson. 1991. Amphibians and Reptiles of the West Estrada, A.R., J. Novo, and L.V. Moreno. 1987. Datos sobre una puesta de Ameiva Indies: Descriptions, Distributions, and Natural History. University of Florida auberi Cocteau (Sauria: Teiidae). Miscelanea Zoológica 30:2–3. Press, Gainesville. Fong, A., D. Maceira, W.S. Alverson, and J.M. Shopland (eds.). 2005. Cuba, Simmons, J.E. 1975. The female reproductive cycle of the teiid lizard Ameiva Siboney-Juticí. Rapid Biological Inventories Report 10. The Field Museum, ameiva petersii. Herpetologica 31:279–282. Chicago, Illinois. Simmons, P.M., B.T. Greene, K.E. Williamson, R. Powell, and J.S. Parmerlee, Gali, F. and O.H. Garrido. 1986. Two new subspecies of Ameiva auberi (Reptilia: Jr. 2005. Ecological interactions within a lizard community on Grenada. Teiidae) from Cuba. Caribbean Journal of Science 22:165–173. Herpetologica 61:124–134. Henderson, R.W. and R. Powell. 2009. Natural History of West Indian Reptiles and Smith, R.E. 1968. Studies of reproduction in Costa Rican Ameiva festiva and Amphibians. University Press of Florida, Gainesville. Ameiva quadrilineata (Sauria: Teiidae). Copeia 1968:236–239. Hirth, H.F. 1963. The ecology of two lizards on a tropical beach. Ecological Somma, C.A. and G.R. Brooks. 1976. Reproduction in Anolis oculatus, Ameiva Monographs 33:83–112. fuscata, and Mabuya mabouya from Dominica. Copeia 1976:249–256. Hower, L.M. and S.B. Hedges, S.B. 2003. Molecular phylogeny and biogeography Sproston, A.L., R.E. Glor, L.M. Hartley, E.J. Censky, R. Powell, and J.S. of West Indian teiid lizards of the genus Ameiva. Caribbean Journal of Science Parmerlee, Jr. 1999. Niche differences among three sympatric species of 39:298–306. Ameiva (Reptilia: Teiidae) on Hispaniola. Journal of Herpetology 33:131–136. Kerr, A.M., R. Powell, and J.S. Parmerlee, Jr. 2005. Ameiva erythrocephala (Teiidae) Wolcott, G.N. 1923. The food of Porto Rican lizards. Journal of the Department of on Sint Eustatius, Netherlands Antilles: Baseline data on a small population in Agriculture of Porto Rico 7:5–38.